Reproduction

Generally, most mammalian species are either polygynous (one male mates with multiple females) or promiscuous (both males and females have multiple mates in a given reproductive season). Because females incur such high costs during gestation and lactation, it is often the case that male mammals can produce many more offspring in a mating season than can females. As a consequence, the most common mating system in mammals is polygyny, with relatively few males fertilizing multiple females and many males fertilizing none. This scenario sets the stage for intense male-male competition in many species, and also the potential for females to be choosy when it comes to which males will sire her offspring. As a consequence of the choices females make and the effort males put into acquiring matings, many mammals have complex behaviors and morphologies associated with reproduction. Many mammal groups are marked by sexual dimorphism as a result of selection for males that can better compete for access to females. (Apfelbach, 1990; Nowak, 1991; Vaughan, Ryan, and Czaplewski, 2000)

About 3 percent of mammalian species are monogamous, with males only mating with a single female each season. In these cases, males provide at least some care to their offspring. Often, mating systems may vary within species depending upon local environmental conditions. For example, when resources are low, males may mate with only a single female and provide care for the young. When resources are abundant, the mother may be able to care for young on her own and males will attempt to sire offspring with multiple females. (Apfelbach, 1990; Vaughan, Ryan, and Czaplewski, 2000)

Other mating systems such as polyandry can also be found among mammals. Some species (e.g. Callithrix jacchus and Panthera leo) display cooperative breeding, in which groups of females, and sometimes males, share the care of young from one or more females. Heterocephalus glaber have a unique mating system among mammals. Like social insects (Hymenoptera and Isoptera), naked mole rats are eusocial, with a queen female mating with several males and bearing all of the young in the colony. Other colony members assist in the care of her offspring and do not reproduce themselves. (Apfelbach, 1990; Keil and Sachser, 1998; Lazaro-Perea et al., 2000; Stockley, 2003; Vaughan, Ryan, and Czaplewski, 2000)

Many mammals are seasonal breeders, with environmental stimuli such as day length, resource intake and temperature dictating when mating occurs. Females of some species store sperm until conditions are favorable, after which their eggs are fertilized. In other mammals, eggs may be fertilized shortly after copulation, but implantation of the embryo into the uterine lining may be delayed (“delayed implantation”). A third form of delayed gestation is "delayed development", in which development of the embryo may be arrested for some time. Seasonal breeding and delays in fertilzation, implantation, or development are all reproductive strategies that help mammals coordinate the birth of offspring with favorable environmental conditions to increase the chances of offspring survival. (Vaughan, Ryan, and Czaplewski, 2000)

Some mammals give birth to many altricial young in each bout of reproduction. Despite being born in a relatively underdeveloped state, young of this type tend to reach maturity relatively quickly, soon producing many altricial young of their own. Mortality in these species tends to be high and average lifespans are generally short. Many species that exemplify this type of life history strategy can be found among the Rodentia and Insectivora. At the other end of the life history spectrum, many mammals give birth to one or a few precocial young in each bout of reproduction. These species tend to live in stable environments where competition for resources is a key to survival and reproductive success. The strategy for these species is to invest energy and resources in a few, highly developed offspring that will grow to be good competitors. Cetacea, Primates and Artiodactyla are examples of orders that follow this general pattern. (Vaughan, Ryan, and Czaplewski, 2000; Wilson and Reeder, 1993)

Among mammals, many reproductive strategies can be observed, and the patterns listed above are the extremes of a continuum encompassing this variation. Environmental factors, as well as physiological and historical constraints all contribute to the pattern of reproduction found in any population or species. Differences in these factors among species have led to the diversity of life history traits among mammals. (Vaughan, Ryan, and Czaplewski, 2000; Wilson and Reeder, 1993)

A fundamental component of mammalian evolution, behavior, and life history is the extended care females must give to their offspring. Investment begins even before a female's eggs become fertilized. All female mammals undergo some form of estrus cycle in which eggs develop and become ready for potential fertilization. Hormones regulate changes in various aspects of female physiology throughout the cycle (e.g., the thickening of the uterine lining) and prepare the female for possible fertilization and gestation. Once fertilization occurs, females nurture their embryos in one of three ways--either by attending eggs that are laid externally (Prototheria), nursing highly altricial young (often within a pouch, or "marsupium"; Metatheria), or by nourishing the developing embryos with a placenta that is attached directly to the uterine wall for a long gestation period (Eutheria). Gestation in eutherians is metabolically expensive. The costs incurred during gestation depend upon the number of offspring in a litter and the degree of development each embryo undergoes. (Vaughan, Ryan, and Czaplewski, 2000)

Once the young are born (or hatch, in the case of monotremes) females feed their newborn young with milk, a substance rich in fats and protein. Because females must produce this high-energy substance, lactation is far more energetically expensive than gestation. Once mammals are born they must maintain their own body temperatures, no longer being able to depend on their mother for thermoregulation, as was the case during pregnancy. Lactating females must provide enough milk for their offspring to maintain their body temperatures as well as to grow and develop. In addition to feeding their young, females must protect them from predators. In some species, young remain with their mothers even beyond lactation for an extended period of behavioral development and learning. (Vaughan, Ryan, and Czaplewski, 2000)

Depending upon the species and environmental conditions, male mammals may either provide no care, or may invest some or a great deal of care to their offspring. Care by males often involves defending a territory, resources, or the offspring themselves. Males may also provision females and young with food. (Apfelbach, 1990)

Mammalian young are often born in an altricial state, needing extensive care and protection for a period after birth. Most mammals make use of a den or nest for the protection of their young. Some mammals, however, are born well-developed and are able to locomote on their own soon after birth. Most notable in this regard are Artiodactyla such as Connochaetes or Giraffa camelopardalis. Cetacea young must also swim on their own shortly after birth. (Vaughan, Ryan, and Czaplewski, 2000)